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Home , Dutch elm disease, Elm, Elm leaf beetle, Ulmus glabra, Verticillium

Dutch Disease AND DISEASE-RESISTANT (DED) has had a devastating impact on the urban of North America. The American elm () was at one time the most extensively planted shade in the . Unfortunately, all elm native to North America, especially Ameri- can elm, are susceptible to DED. The disease was first described in the in 1919. By the early 1930’s the first outbreaks of DED in North America had occurred in Cleveland and Cincinnati, . The first reported cases in Iowa were in Lee and Scott Counties in 1956. Since that time, DED has been detected in all 99 counties, and approxi- mately 95 percent of Iowa’s urban American elms have been killed.

SUL-4 | January 1999 2

by Cassandra Biggerstaff, graduate student, Department of Pathology; Jeffery K. Iles, assistant professor, Department of Horticulture; and Mark L. Gleason, professor, Department of , Iowa State University

DED has destroyed a similar percentage of urban To mail samples directly to the Plant Disease Clinic, elms throughout the United States. Millions of native wrap them in dry newspaper, place in an open plastic elms in the countryside also have died from the disease, bag, and then in a box. Include your name and ad- and continue to die every year. Threatened elm dress, and specify that you want the branches tested for trees can be preserved only when an intensive, coop- DED. Address the samples to the Plant Disease Clinic, erative effort is made to control Dutch elm disease. 351 Bessey Hall, Iowa State University, Ames, IA 50011. A hopeful development in the war against DED is A small fee is assessed for this service. the commercial release of a number of elm and species that possess genetic resistance or tolerance to Causal Agent Dutch elm disease is caused by the disease. The characteristics of these elms will be the fungi ulmi and Ophiostoma novo-ulmi. discussed in the second part of this bulletin. 0. ulmi is the original species introduced to the United States and . O. novo-ulmi is a more aggressive Symptoms The first symptom of Dutch elm species that has become more prevalent in recent de- disease is wilting on one or a few branches in the cades, virtually replacing O. ulmi in Iowa. Ophiostoma upper canopy of the tree (Figures 1 and 2). Affected grows in the (water-conducting tissue) of elms. leaves usually turn yellow, then brown, and fall from In an attempt to fight invasion by the , an in- the tree prematurely. Sometimes the leaves dry out very fected tree will produce gums and cell outgrowths that quickly and turn dull green, remaining on the tree for block its own xylem vessels. Blockage of a tree’s water weeks or months before falling. Once a branch has lost supply causes the characteristic wilting symptoms. its leaves, it usually dies quickly. If a tree infected in one season survives to the next season, it will produce smaller-than-normal leaves during spring growth. De- 1 pending on when and how the tree was infected, its branches may die progressively, or the entire tree may Wilting caused by DED has progressed into several die within a few weeks of infection. branches of this tree. Elm leaves also can wilt or turn brown for other 2 reasons, including mechanical injuries, feeding damage by elm (Figure 3), or infection by elm yellows, A single wilted branch in the canopy of an a common northeastern U.S. disease caused by a type American elm is an early symptom of DED of bacterium called a . For this reason, infection that has been spread by elm beetles. you need to look closely to determine if Dutch elm dis- 3 ease is causing the wilting. When the bark is stripped from recently wilted branches of DED-infected trees, Damage to leaves caused by elm leaf beetles is brown streaks are usually present in the outer sapwood unrelated to DED but is sometimes mistaken for (Figure 4). wilting symptoms of this disease. Laboratory testing of branch segments is the best 4 way to confirm whether a wilting elm tree has DED. In Iowa, samples can be submitted to your local county Discoloration of the xylem (water-conducting Extension office or mailed directly to the Plant Disease tissue) caused by DED infection often results in a Clinic at Iowa State University. Samples for testing dark brown discoloration of one-year-old xylem in should be branch segments 1/4 to 1 inch in diameter a ring pattern (center branch) or dark brown and 6 to 12 inches in length, collected from recently streaks in xylem vessels in the outer sapwood wilted areas of the tree. At least four branch segments (branch at right). The branch at left displays the that show brown streaking should be collected. The normal cream color of healthy . DED fungus cannot be isolated from dry, dead branches. 1 2 3 4 4

Disease Transmission The fungus is trans- within the xylem to any other elm tree which is root- mitted from infected to healthy trees either by elm bark grafted to the infected tree. This is the reason neighbor- beetles or by root grafts formed between neighboring hoods with colonnades of adjacent elms lining streets trees. lost many trees very rapidly (Figures 10 and 11). It has Bark transmission. Transmission by elm bark been estimated that more than 90 percent of urban elms beetles is a key component of the DED disease cycle. killed in DED epidemics became infected through root Two species of beetles spread the fungus from infected grafts. to healthy elms: the European elm ( multistriatus) (Figures 5 and 6) and the native elm bark Control Strategies . Prompt, thor- beetle (Hylurgopinus rufipes). These two beetles are simi- ough sanitation is vital for controlling Dutch elm dis- lar in size (approximately 1/8 inch long) and appear- ease. DED sanitation programs can be successful only ance, but they have different habits and life cycles. when they are applied to the entire community, includ- European elm bark beetles, the more common of ing trees in parks, fencerows, cemeteries, and other sites the two beetle species in Iowa, survive the winter as as well as in residential areas. Removing sources of the larvae in recently killed elm wood. The DED fungi grow DED fungus and breeding grounds for the beetles is into the beetles’ brood galleries (tunnels in the inner essential. Any tree that has died due to DED should be bark where eggs are laid). The sticky spores of the DED cut down and destroyed promptly. Branches pruned fungi (Figure 7) will adhere to adult beetles as they emerge from infected trees also should be destroyed. All in- in late April through June. The beetles then fly to a live fected wood should be burned, buried, or chipped. Do elm tree and bore into small branch crotches to feed. not store firewood from DED-infected elms with the bark Germinating spores on the beetle’s body enter the vas- attached, because logs with attached bark provide breed- cular system of the tree through feeding wounds. After ing sites for beetles. Available evidence indicates that maturing, adult females seek out sick or dead elm wood in which to lay eggs. The female lays her eggs in brood galleries tunneled parallel with the in the inner bark. Larvae hatched from these eggs bore tun- nels from the main brood gallery into the surrounding 5 inner bark (Figure 8). Native elm bark beetles overwinter as larvae in dead Size comparison of elm bark beetles with the end wood or as adults in the lower trunks of healthy elms. of a pencil. Overwintering adults emerge in the spring and tunnel 6 into the inner bark of branches two to four inches in diameter. As the weather turns warmer, the beetles leave Close-up of a European elm bark beetle, the their branches to find suitable dead wood in which to major vector of DED in Iowa. lay their eggs. The brood galleries of the native bark 7 beetle are bored perpendicularly to the grain of the wood, and developing larvae bore away from the gallery in the Sticky spores of the DED fungi, produced in the direction of wood grain. Larvae of the native elm bark bark beetle galleries of an infected elm, adhere to beetle mature and emerge as adults in late July and Au- the bodies of elm bark beetles before the beetles gust. emerge from these trees and fly to healthy trees. Root Graft Transmission. The second way DED fungi 8 can enter healthy elm trees is through root grafts (Figure 9). The root systems of elms whose trunks are within Larvae (white semicircles) are visible in tunnels approximately 50 feet of one another tend to fuse to- they bored through the inner bark of an elm tree gether, producing a continuous vascular network be- from the brood gallery (large vertical tunnel), tween trees. If one of the trees in the network becomes where eggs that hatched into the larvae were laid. infected with Ophiostoma, the fungus can move rapidly 5 6 7 8 6

wood chips made from DED-infected trees pose no risk containing concentrated . These capsules can of transmitting the disease. be pressurized by hand after their injection tees are Disrupting Root Grafts. When the trunk of a DED- tapped into holes in the root flare. Although it is legal infected elm is within 50 feet of the trunks of other sus- for homeowners to apply injection treatments to their ceptible elms, root grafts can be disrupted to block the own trees, it is strongly recommended that injections spread of the disease. Determine the line of root graft be done by a reputable professional arborist who also is disruption between adjacent trees by marking a line or a licensed pesticide applicator. an arc equidistant between the two trees. This is the Fungicide injections are costly. A single mature line along which trenching should be done, or soil fu- elm will often cost hundreds or even thousands of dol- migants placed. Trenching can be done with a chain lars to inject. The protective effect of fungicide injec- trencher, a vibratory plow, or a sharp spade. The trench tions may last one to three years, so repeat injections should be at least four feet deep. Be sure to contact lo- may be needed. Therefore, only very valuable trees are cal authorities about the possibility of buried utilities treated with . Fungicide treatments alone before digging (In Iowa, call 1-800-292-8989). In ar- cannot save an elm tree from succumbing to DED, but eas where trenching is not an option due to sidewalks, along with prompt, thorough sanitation they can help fences, or buried utilities, chemical fumigants can be an save high-value trees that would otherwise be killed. effective way to disrupt root grafts. Holes two feet deep and close to an inch in diameter should be dug along the equidistant line every six to twelve inches. A prop- erly labeled fumigant such as Vapam can be poured into these holes, which are then sealed. When working around sidewalks and driveways, holes should be angled underneath these features to more effectively distribute the fumigant. Be sure to follow label directions care- 9 fully. Excavated roots of two adjacent elms showing Fungicide injection. Fungicide injections have between the roots. Root grafting joins been used to defend against DED for more than 50 years. together the vascular systems of adjacent trees, Fungicide injections have been most successful in con- allowing DED fungi to pass through grafts from an trolling Dutch elm disease when used as a preventive infected tree into its neighbor. treatment on uninfected trees. Success in treating trees with crown symptoms is much less certain. Trees with 10 more than 10 percent of the crown infected are unlikely Rows of adjacent American elms arching over a city to be rescued by fungicide injection. street provided a beautiful colonnade effect. This Several methods are used for injecting fungicides distinctive landscape feature was common before into elms. One method is to pump a dilute solution of DED epidemics killed most such plantings. fungicide into a tree through holes drilled in the root flare. The root flare, the zone where the trunk meets 11 the roots, is exposed by excavating soil around the trunk Extensive colonnades of American elms were killed to a depth of 6 to 12 inches. Small (7/32- to 9/32-inch- rapidly during DED epidemics. Most trees in such diameter) holes are drilled less than 1 inch deep into colonnades were killed when the DED fungi spread the root flares around the entire circumference of the between neighboring trees through root grafts. tree at specified spacing. After tapered injection tees are tapped into the holes, the tees are connected to each 12 other and to a pump with plastic tubing (Figure 12). The root flare of this American elm has been The fungicide is pumped into the tree through the mani- excavated to permit installation of plastic tees and fold of plastic tubing and injection tees. An alternative tubing for injection of fungicides injection technique involves the use of plastic capsules to protect against DED. 9 10 11 12 8 DED-resistant Elms Fungicides and sanitation have had only occasional suc- has demonstrated tolerance to DED, but may be quite cess in deterring DED. Researchers have therefore redi- susceptible to elm yellows (a relatively rare disease in rected their efforts to developing DED-tolerant or DED- the Midwest). resistant elms. European and Asiatic elms have played a The combination of environmental adaptability, significant role in the breeding, selection, and DED tolerance, and the highly prized vase-shaped crown release efforts, but considerable emphasis also has been form will make these selections extremely popular in given to breeding and selecting American elms. The goal the future. However, it will probably take a number of of finding or creating a DED-tolerant American elm was years before these trees become commonplace in nurs- viewed skeptically in the past, but hopes have been re- eries and retail garden centers. It also is important to vived with the release of several cultivars showing high remember that none of the new American elm cultivars levels of DED tolerance. are completely immune to DED, and that DED toler- The new American elm cultivars, ‘Valley Forge’ ance of mature trees has not yet been evaluated. (Figure 13) and ‘New Harmony’ (Figure 14), are prod- Homeowners and professional arborists also should ucts of the U.S. National Arboretum tree genetics pro- consider the many commercially available elms gram that have demonstrated high levels of tolerance to for augmenting and diversifying their plantings. These both O. novo-ulmi and O. ulmi. Tolerance to DED is char- hybrids are the results of crosses made with species of acterized by reduced wilting and crown dieback after elms from and Europe which have very high levels fungal inoculation. In fact, these trees have the ability to of resistance to DED. Some of the more important in- recover from DED infection even after symptom expres- troductions are described below. sion. AccoladeTM (Ulmus japonica x Ulmus wilsoniana) Of the thousands of American elms screened, ‘Val- (Figure 16)— Also known as Thornhill elm. This hy- ley Forge’ has shown the best tolerance to DED. The brid was released by the Morton Arboretum, Lisle, IL. tree has an upright, arching, broadly vase-shaped branch- ing structure with a full, dense canopy of leaves. Clones from the original parent tree are 26 feet tall with an av- erage crown spread of 30 feet after 12 growing seasons. Summer leaves are green, turning yellow in autumn. The bark is typical of the species, with grayish, flat-topped ridges separated by diamond-shaped fissures. ‘Valley Forge’ is considered hardy in USDA zones 5 (southern Iowa) through 7. 13 While not as disease-tolerant as ‘Valley Forge,’ ‘New ‘Valley Forge,’ a DED-tolerant American elm Harmony’ still ranks in the top three among the thou- cultivar developed at the U.S. National Arboretum. sands of American elms subjected to intensive inocula- tion with the DED fungus. The parent tree of ‘New Har- 14 mony’ displays a broad, vase-shaped crown and has ‘New Harmony,’ a DED-tolerant American elm grown approximately 68 feet tall and 72 feet wide. Leaf cultivar developed at the National Arboretum. and bark characteristics are similar to those of ‘Valley Forge’, but because of its acceptable performance in 15 Minnesota, ‘New Harmony’ is considered hardy in USDA zones 4 through 7 (includes all of Iowa). ‘Independence,’ a DED-tolerant American elm ‘Independence’ (Ulmus americana ‘Moline’ x Ulmus hybrid developed at the University of Wisconsin. americana W 185-21) (Figure 15) is one of six clones 16 that comprise the much-heralded ‘American Liberty’ multiclone variety. Patented by Smalley and Lester (Univ. AccoladeTM, a hybrid elm developed of Wisconsin, Madison), ‘Independence’ develops an at the Morton Arboretum. upright, vase-shaped crown typical of the species and 13 14 15 16 10

The tree displays a handsome vase-shaped canopy, deep moderately vase-shaped crown, resembling a more up- green glossy leaves, and has shown resistance to DED, right American elm. It has shown a high level of resis- elm , and leaf miner, but only moderate toler- tance to DED, high tolerance to elm yellows, and re- ance of urban soil conditions such as high clay content duced susceptibility to the . ‘Patriot’ is and seasonal wetness. It is hardy through USDA zone adapted to a wide variety of soil conditions, grows best 4. Other elms from the Morton Arboretum that show in full sun, and is considered cold hardy through USDA promise include Danada CharmTM, ‘Vanguard,’ and ‘Tri- hardiness zone 4. umph.’ ‘Pioneer’ ( x Ulmus carpinifolia) (Fig- ‘Cathedral’ ( x Ulmus japonica) (Fig- ure 22)— ‘Pioneer’ is a vigorous, fast-growing USDA se- ure 17) — One of several excellent cultivars developed lection with large, dark green leaves and a globe-shaped at the University of Wisconsin, ‘Cathedral’ has demon- crown. Hardy in USDA zones 5 through 8, it has proven strated good tolerance to DED, experiencing only branch resistant to DED and elm yellows, but elm leaf beetle tip injury when infected with the fungus. The tree has a feeding may be a problem. Because of its broad, spread- broad vase shape, medium to light green leaves in sum- ing habit, ‘Pioneer’ is best suited to spacious grounds mer, and yellow fall foliage. In addition, ‘Cathedral’ is like those found in parks, golf courses, and large com- highly tolerant to wilt and is resistant to at- mercial properties. tack by the elm leaf miner. It is reliably hardy through ‘Prospector’ (Ulmus wilsoniana) (Figure 23) — This USDA zone 4. seedling selection was released in 1990 by the USDA. ‘Frontier’ (Ulmus carpinifolia x ) ‘Prospector’ elm has an American elm-like vase-shaped (Figure 18) — Released in 1990 by the USDA, ‘Frontier’ crown but its branches become pendulous at a much has demonstrated a high degree of resistance to DED, lower height. Newly expanding leaves are orange-red, moderate resistance to elm leaf beetle, and high toler- but gradually darken to green, finally turning yellow in ance to the phytoplasma-caused elm yellows. autumn. ‘Prospector’ is resistant to DED, tolerant to elm Emerging leaves in spring are red, gradually chang- yellows, resistant to elm leaf beetle, and is considered ing to yellow-green in summer, finally turning red-purple adaptable in USDA hardiness zones 4 through 7. in autumn. Although ‘Frontier’ becomes pyramdial in- stead of vase-shaped as it matures, it still should make a desirable street, park, landscape, or highway tree. Be- cause it has sustained some low-temperature injury in Minnesota, ‘Frontier’ is considered reliably hardy only through USDA zone 5 (southern Iowa). ‘Homestead’ (Ulmus pumila x complex hybrid from the Netherlands elm breeding program) (Figure 19) — Another USDA release, ‘Homestead’ has a symmetrical, somewhat pyramidal crown that becomes arching as the 17 tree ages. Its dark green summer leaves turn golden- yellow in fall and the growth rate is reportedly rapid. ‘Cathedral,’ a hybrid elm developed ‘Homestead’, best used in USDA hardiness zones 5 at the University of Wisconsin. through 8, is considered highly resistant to DED, but is 18 susceptible to elm leaf beetle. ‘New Horizon’ (Ulmus japonica x Ulmus pumila) ‘Frontier,’ a hybrid elm developed by the USDA. (Figure 20) — This hybrid has excellent resistance to 19 DED and elm leaf miner and high tolerance to Verticil- ‘ lium wilt. It has an upright habit, strong branch struc- ‘Homestead,’ a hybrid elm released by the USDA. ture, and a dense crown with dark green leaves. ‘New 20 Horizon’ is hardy through USDA zone 4. ‘ ‘Patriot’ (Ulmus ‘Urban’ x Ulmus wilsoniana ‘Pros- ‘New Horizon,’ a hybrid elm cultivar released pector’) (Figure. 21) — Developed at the U.S. National by the University of Wisconsin. Arboretum and released by the USDA, ‘Patriot’ has a 17 18 19 20 12

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‘Patriot,’ a hybrid elm developed by the ‘Regal’ (Ulmus ‘Commelin’ x Ulmus ‘Hoersholmiensis’) National Arboretum and released by the USDA (Figure 24) — Selected at the University of Wisconsin, ‘Re- gal’ develops a strong central leader with an upright or 22 columnar growth habit when young, becoming more ovate with age. Leaves are dark green in summer, show no ap- ‘Pioneer,’ a hybrid elm released by the USDA. preciable fall coloration, and because they are rather 23 sparsely borne, cast a honeylocust-like light shade that makes possible the successful culture of turfgrass in the ‘Prospector,’ a seedling elm released by the USDA. vicinity of the tree. ‘Regal’ is considered highly resistant to 24 DED and , and is hardy in USDA zones 4 through 7. ‘Regal,’ a hybrid elm released by the Over the past 30 to 40 years, plant breeders have spent University of Wisconsin. considerable time, effort, and resources searching for elms having resistance to DED and other pests, and possessing the graceful crown architecture of the revered American Cover photo by Cameron Davidson; elm. The recent release of Ulmus americana ‘Valley Forge’ Other photos by E.B. Smalley, University of Wisconsin; and ‘New Harmony’ and the improved availability of many D. Townsend, U.S. National Arboretum; University of excellent hybrid elms are certain to spark renewed inter- Illinois; D.R. Lewis, Iowa State University; and est for elms of all kinds. However, the sad experience of A.H. Epstein, Iowa State University the DED epidemic has taught some hard lessons about using elms in urban sites. It is clearly important to avoid [D] File: Pest Management 5 4/01 colonnade-type plantings of elms, since adjacent trees will . . . and justice for all The U.S. Department of Agriculture (USDA) prohibits discrimination root-graft and potentially spread DED. The new elm cul- in all its programs and activities on the basis of race, color, national origin, gender, religion, age, disability, political beliefs, sexual tivars and species are best used as specimen trees rather orientation, and marital or family status. (Not all prohibited bases than as large-scale monocultures as in the past. With ge- apply to all programs.) Many materials can be made available in alternative formats for ADA clients. To file a complaint of discrimina- netic tolerance and resistance, together with judicious tion, write USDA, Office of Civil Rights, Room 326-W, Whitten planting practices, the elm may be ready for a major come- Building, 14th and Independence , SW, Washington, DC 20250-9410 or call 202-720-5964. back in Iowa .␣ Issued in furtherance of Cooperative Extension work, Acts of May 8 and June 30, 1914, in cooperation with the U.S. Department of Agriculture. Stanley R. Johnson, director, Cooperative Extension Service, Iowa State University of Science and Technology, Ames, Iowa.